Journal of Oral Rehabilitation 2006 33; 262273

Review Article

Sensory-motor function of human periodontal

mechanoreceptors*
M. TRULSSON

Institute of Odontology, Karolinska Institutet, Huddinge, Sweden

Natural teeth are equipped with periodontal mechanoreceptors that signal information
about tooth loads. In the present review, the basic
force-encoding properties of human periodontal
receptors will be presented along with a discussion
about their likely functional role in the control of
human mastication. Microneurographic recordings
from single nerve fibres reveal that human periodontal receptors adapt slowly to maintained tooth
loads. Most receptors are broadly tuned to the
direction of force application, and about half respond to forces applied to more than one tooth.
Populations of periodontal receptors, nevertheless,
reliably encode information about both the teeth
stimulated, and the direction of forces applied to
the individual teeth. Information about the magnitude of tooth loads is made available in the mean
firing rate response of periodontal receptors. Most
SUMMARY

Introduction
To control oral motor behaviours such as biting, chewing, speech and oral manipulation, the brain relies on
information from sense organs in the orofacial structures (13). Natural teeth are equipped with extremely
sensitive tactile sensors periodontal mechanoreceptors
(2, 4, 5). These sensors provide information about tooth
loads and are located among the collagen fibres in the
ligaments that attach the root of the tooth to the alveolar
bone. If we are to understand how periodontal receptors
contribute to the control of oral motor behaviours, a
description of the information they carry to the brain
during these behaviours is essential. With this know*Based on the Journal of Oral Rehabilitation Summer School 2005 in
Bevagna, Italy. Kindly sponsored by Blackwell Munksgaard and Nobel
Biocare.
2006 Blackwell Publishing Ltd

receptors exhibit a markedly curved relationship

between discharge rate and force amplitude, featuring the highest sensitivity to changes in tooth load at
very low force levels (below 1 N for anterior teeth
and 4 N for posterior teeth). Thus, periodontal
receptors efficiently encode tooth load when subjects contact and gently manipulate food using the
teeth. It is demonstrated that signals from periodontal receptors are used in the fine motor control
of the jaw and it is clear from studies of various
patient groups (e.g. patients with dental implants)
that important sensory-motor functions are lost or
impaired when these receptors are removed during
the extraction of teeth.
KEYWORDS: mechanoreceptor, microneurography,
sensory-motor control, tooth, implant, mastication
Accepted for publication 27 January 2006

ledge, it is possible to formulate and test hypotheses on

the likely functional role of periodontal receptors. Solid
knowledge on how the brain utilizes sensory signals to
regulate oral motor behaviours is a prerequisite for
understanding the functional consequences of many
clinical treatments. For example, when a natural tooth
is replaced by a dental implant the periodontal ligament
disappears and the information from periodontal receptors about tooth loads is no longer available for the
regulation of oral motor functions. In this review, the
basic force-encoding properties of human periodontal
receptors including a description of their signalling
during biting and chewing will be presented and
discussed in the light of their likely functional role in
the control of oral motor actions. The functional
consequences of replacing natural teeth with oral
implants will also be commented on.
doi: 10.1111/j.1365-2842.2006.01629.x

SENSORY-MOTOR FUNCTION OF HUMAN PERIODONTAL MECHANORECEPTORS

Recording signals from human periodontal

receptors
Nerve signals were recorded from single periodontal
afferents innervating anterior and posterior teeth in
healthy adult volunteers. The neural data were
obtained by the microneurographic technique developed by Vallbo and Hagbarth (6). The inferior alveolar
nerve was approached through the oral cavity and
impaled by a tungsten needle electrode near its
entrance at the mandibular foramen (Fig. 1a) (7, 8).
Manual handling of the electrode was guided by
landmarks used in dental practice when blocking the
mandibular nerve. Once the tip of the electrode was
located in a nerve fascicle, its position was carefully
adjusted until impulse responses from a single afferent
were obtained. The electrode was then left undisturbed
in this recording position, supported only by the
surrounding tissues.
To activate periodontal receptors during the search
for single-unit responses, light mechanical stimuli
(<5 N) were applied manually to the crowns of the
teeth. The search strategy adopted in these experiments
was optimized for low-threshold mechanoreceptive
afferents. Thus, other types of afferents that may
contribute to dental mechanosensitivity (e.g. highthreshold periodontal afferents and intra-dental afferents) were not studied and will not be further discussed
(e.g. see Ref. 9).
When stable recordings were obtained from single
periodontal afferents, controlled forces were applied to
small nylon cubes cemented to the incisal edges or
occlusal surfaces on individual teeth. The neural
discharge was recorded in response to forces applied
in four directions in the horizontal plane (lingual,
facial, mesial, distal) and in the two axial directions
(up and down; Fig. 1b). Recordings from the inferior
alveolar nerve revealed that human periodontal
mechanoreceptors discharge continuously during sustained loading of the teeth. However, consistent with
animal studies (see Refs 4, 5), it was found that the
receptors slow-adapting nature might not be recognized if the stimuli are limited to directions of
application that activate the receptors poorly. Like
the slow-adapting type II (SA II) mechanoreceptors in
the skin, most human periodontal mechanoreceptors
(about 70%) are active spontaneously and discharge
regularly in response to forces applied to the teeth
(Fig. 2a).

(a)

(b)

Fig. 1. (a)Recording from periodontal mechanoreceptors in the

inferior alveolar nerve in man. Forces are applied to the central
incisor by the use of a hand-held probe equipped with force
transducers (FT) for continuous force measurement. Note the small
nylon sphere at the end of the probe and the nylon cubes attached
to the teeth. (b) A tooth is shown in the horizontal plane (left) and
in a vertical plane (right). Each of the six directions of stimulation
are represented by the arrows: Fa, Me, Li, Di, Do and Up refer to
facial, mesial, lingual, distal, downward and upward, respectively.
The forces were applied to each of the five free faces on the nylon
cube cemented to the tooth just above its edge. The probe slipped
off the cube if the force was not applied perpendicular to the faces,
ensuring that data were collected only in the desired direction of
force application. The upward force (Up) was applied with a nylon
loop (not shown) [Reprinted from Progress in Neurobiology, Vol.
49 Trulsson M, Johansson RS. Encoding of tooth loads by human
periodontal afferents and their role in jaw motor control pp 267
284, Copyright 1996, with permission from Elsevier].

Receptor encoding of spatial aspects of

tooth loads
Typically, periodontal mechanoreceptors are broadly
tuned to the direction of tooth loading (Fig. 2). That is,
the afferents normally respond to forces applied to the
receptor-bearing tooth in two to four of the six test
directions (8, 10). These receptive field properties are in
concordance with those observed in the cat, but are in
contrast to those in the dog, which appeared to have
narrower receptive fields (1113). As a result of the
broad directional tuning, an individual periodontal

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M. TRULSSON

Fig. 2. Directional sensitivity of human periodontal receptors. (ab) Responses of a single periodontal receptor to forces applied in six
directions to the lower central incisor (the receptor-bearing tooth). (a) Examples of nerve recordings with the corresponding force records
above each. (bf) Vectorial representation of the responses for five afferents to sustained force application in the horizontal (left) and axial
(right) directions. (b illustrates the afferent shown in a). The length of each vector is proportional to the mean discharge rate evoked in
each direction. The spontaneous discharge rate is represented by the circle with the radius indicating its intensity. Vectors longer and
shorter than this radius illustrate increased and decreased firing, respectively. The thick arrow represents an estimate of the most efficient
excitatory stimulus direction in the horizontal plane, i.e. the calculated preferred direction [Reprinted from the Journal of Physiology . Directional sensitivity of human periodontal mechanoreceptive afferents to
London, Vol. 447, Trulsson M, Johansson RS, Olsson KA
forces applied to the teeth, pp 373389, Copyright 1992, with permission from Blackwell Publishing].

mechanoreceptor provides ambiguous information

about the direction of force applied to the tooth.
However, analyses based on vector calculations show
that information about the precise direction of force is
represented reliably in the activity from small populations of periodontal mechanoreceptors (8).
The receptive field of human periodontal mechanoreceptors often extends beyond a single tooth. About
half of the periodontal mechanoreceptors respond to
loading of groups of adjacent teeth, typically two to four
teeth (10, 14). Each afferent always exhibits the highest
response rates to stimulation of one particular tooth,
the so-called receptor-bearing tooth, with a gradual and
rather sharp decline in responsiveness to loads applied
to the adjacent teeth (Fig. 3). Moreover, the preferred
directions of the different teeth strongly suggest that
multi-tooth receptive fields result from mechanical
coupling between neighbouring teeth by interdental
contacts and trans-septal collagen fibres, rather than by
branching of axons to more than one tooth.
The multiple-tooth characteristics of the receptive
fields imply that during mastication not only are the
periodontal afferents that innervate the tooth to which
a force is directly applied activated, but so are those that

innervate the adjacent teeth. However, considering the

rather sharp decline in both number of activated
afferents and response intensities from the receptorbearing tooth to the adjacent teeth, we hypothesized
that the population of human periodontal afferents
accurately encodes the location of the tooth directly
loaded. By the use of artificial neural networks it was
shown that the population response in human periodontal afferents indeed contains detailed information
about both the direction of applied forces and the tooth
loaded (15). Thus, the existence of multiple-tooth receptive fields does not downgrade the ability of the central
nervous system to uniquely locate a force stimulus
striking a particular tooth. Overlapping peripheral
receptive fields are characteristic of all sensory systems
and have been shown to improve, rather than degrade,
their spatial localization and resolution (e.g. 16).
Periodontal receptors of anterior and posterior teeth
do indeed demonstrate similarities but there are also
some important differences (10, 17). The use of vector
calculations to identify differences in the periodontal
receptors at different types of teeth is illustrated in
Fig. 4. Based on the receptors response to forces
applied in each of the six test directions, its single

2006 Blackwell Publishing Ltd, Journal of Oral Rehabilitation 33; 262273

SENSORY-MOTOR FUNCTION OF HUMAN PERIODONTAL MECHANORECEPTORS

Fig. 3. Receptive field of a single periodontal mechanoreceptor

showing steady-state responses to maintained mechanical stimulation of five teeth, illustrated from a horizontal view (a) and a
facial view (b). The vectors illustrate the receptors response to a
tooth load of 250 mN. The tested teeth: c1 contralateral central
incisor, 1 central incisor, 2 lateral incisor, 3 canine,
4 first premolar. RBT, receptor-bearing tooth [Reprinted from
the Journal of Neurophysiology, Vol. 69, Trulsson M. Multiple
tooth receptive fields of single human periodontal mechano
receptive afferents, pp 474481, Copyright 1993, with permission
from the American Physiological Society].

preferred direction was calculated by vector summation (see the thick arrows in Fig. 2bf). The vectors in
Fig. 4 represent the preferred directions for all periodontal receptors that have been studied in man. Each
cluster of arrows represents the vectors calculated for all
of the receptors that were examined in all teeth of that
type (anterior teeth, first premolars, the second premolars and the first molars).
Note that the number of vectors decreases from the
anterior teeth to the molar, indicating a decreasing
number of receptors in the periodontal ligaments
distally along the dental arch. This finding is supported
by several histological studies (1820) and attests to the
importance of a well-developed mechanoreceptive
innervation of the anterior part of the mouth. When
food is taken into the mouth, the morsel will be split
into smaller pieces by the front teeth before it is moved
to posterior parts of the dentition. Sometimes, the front

Fig. 4. Preferred directions of individual receptors of different

teeth projected to the horizontal plane and two vertical planes,
respectively [Reprinted from the Journal of Physiology-London,
. Directional
Vol. 447, Trulsson M, Johansson RS, Olsson KA
sensitivity of human periodontal mechanoreceptive afferents to
forces applied to the teeth, pp 373389, Copyright 1992, with
permission from Blackwell Publishing].

teeth are even used as a third hand in manipulative

tasks, or as a precision cutting tool. Proper execution of
such tasks relies heavily on sensory information as
those involving execution of comparable manipulative
tasks performed by the densely innervated fingertips of
the hand (21).
In the horizontal plane, the preferred directions of
the receptors supplying the anterior teeth and the
premolars are quite evenly distributed around the
circumference of the tooth (Fig. 4). The receptors
supplying the molars, however, have a clear preference
for the distal-lingual direction. In the vertical plane,
there is a preference for downward-directed forces, but
with fewer receptors responding preferentially in this
direction with distance along the arch.
The shift from a high sensitivity to most directions at
the anterior teeth to the distal-lingual direction at the
molars meets the functional demands of the anterior
versus posterior teeth. When the anterior teeth are
manipulating food morsels and splitting them into
pieces in the initial stages of food intake, forces are

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M. TRULSSON
applied to them in all directions. The molars, on the
other hand, grind food substances only during more
forceful chewing. During the final phase of the chewing
cycle, when the lower molars on the working side
approach the intercuspal position from a posterior and
lateral position, they are likely to experience distal and
lingually directed forces upon contact with the opposing upper molar teeth. Studies on tooth displacement
during oral function indicate a tilt of the first molar
during chewing and biting in a lingual direction (22,
23). Given their directional preference for distal-lingual
loading, the mechanoreceptors supplying the lower
molar teeth are well suited to encode information about

the forces that normally act on the posterior teeth

during mastication.

Receptor encoding of intensive aspects of

tooth loads
To study the encoding of intensity of tooth loads by
human periodontal receptors, ramp-and-hold-shaped
force profiles were delivered to the receptor-bearing
tooth in its most responsive direction (Fig. 5a) (17, 24).
The relationships between the magnitude of the sustained force and the steady-state discharge rate for a
group of periodontal receptors supplying the anterior

Fig. 5. (ab) Responses of human periodontal mechanoreceptors to sustained forces of various amplitudes applied to the receptor-bearing
tooth (an incisor or a canine) in the most responsive direction. (a) Examples of force stimulation and nerve recordings of a single afferent
during stimuli of four different amplitudes. (b) Stimulusresponse functions for 19 periodontal receptors. The curves fitted to the data are
defined by the function F/(F + c), where F represents the force, and c the force at which half the estimated maximum discharge rate is
attained. Solid and dashed curves refer to afferents showing saturating stimulusresponse relationships (n 15) and non-saturating
relationships (n 4), respectively. The curve labelled a refers to the same afferent as illustrated in (a). (c) Responses of a saturating and a
non-saturating receptor to an abrupt increment in force superimposed on a sustained force [Reprinted from the Journal of
Neurophysiology, Vol. 72, Trulsson M, Johansson RS. Encoding of amplitude and rate of forces applied to the teeth by human periodontal
mechanorecetive afferents, pp 17341744, Copyright 1994, with permission from the American Physiological Society.
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SENSORY-MOTOR FUNCTION OF HUMAN PERIODONTAL MECHANORECEPTORS

teeth are shown in Fig. 5b. For each periodontal
afferent investigated, the steady-state discharge rate
was approximately proportional to F/(F + c), where F
represents the force, and c the force at which half the
estimated maximum discharge rate is attained. This
transform implies that the discharge rate increases more
or less linearly until F approaches c and then levels off.
Similar stimulusresponse relationships were observed
in less-responsive directions of stimulation, but with
lower discharge rates. These stimulusresponse relationships characterize the static sensitivities of the receptors
to different forces applied to the teeth. The solid curves in
Fig. 5b show that the response to a continuous force is a
markedly downward-curved or hyperbolic relationship
(see solid curves in figure) for most of the human
periodontal mechanoreceptors (about 80%). The slopes
of the curves are steepest between 0 and 1 N, which
indicates that these receptors are most sensitive to
changes in sustained force levels below about 1 N in
magnitude. In contrast to the steeply inclined nature of
the curves below these limits, the curves are almost
horizontal for sustained force levels above the limits. This
indicates that the individual afferents signal the presence
of higher forces but provide the brain with no useful
information as to their magnitudes. That is, the discharge
of each afferent saturates and attains approximately its
highest rate for all force levels above the limit. Saturating
stimulusresponse relationships similar to those of most
human periodontal receptors are also characteristic of
periodontal receptors in animal studies (2528).
In addition to their sensitivity to static forces, periodontal receptors are also sensitive to rapidly changing
forces. Their sensitivity to rapidly changing forces, or
dynamic sensitivity, varies in parallel with their sensitivity to slow changes in static force. This is illustrated
for a saturating receptor on the left of Fig. 5c. After
approximately 1.5 s of exposure to a sustained force,
the tooth was further loaded with a rapid increment in
force. The rapid increase in force evoked an increase in
the steady-state discharge rate that was less than the
addition evoked by the much smaller increment at the
onset of the sustained force. That is, despite the fact that
this force increment was larger and applied at a higher
rate, the receptor discharged more strongly to the
smaller increment in force and lower force rate at the
initial contact with the tooth. This is due to saturating
receptors becoming progressively less sensitive to both
the magnitude and rate of rapid changes in force, as
contact force grows in amplitude.

In contrast to the majority of human periodontal

mechanoreceptors whose responses saturate at very
low forces, a minority (about 20%) do not saturate over
the same range of forces. These non-saturating receptors exhibit stimulusresponse relationships that are
nearly linear (see dotted curves in Fig. 5b). The nonsaturating receptors efficiently encode changes in static
force also at high force levels. As for the saturating
afferents, the dynamic sensitivity of the non-saturating
afferent parallels its static sensitivity. Thus, sensitivity to
rapid changes in force is maintained at high force levels.
This is illustrated for a non-saturating afferent to the
right in Fig. 5c. The strong discharge evoked by the
terminal, rapid increment in force was by far the largest
component of the afferents total response to the
stimulation.
The general shape of the steady-state stimulus
response relationships is similar for periodontal receptors at anterior and posterior teeth (17, 24). For both
posterior (85%) and anterior teeth (about 80%) a
majority of the receptors show a strongly saturating
stimulusresponse relationship. However, compared
with receptors associated with anterior teeth, the
receptors at posterior teeth demonstrate a lower sensitivity at low force levels. Figure 6 illustrates this by the

Fig. 6. Mean value of all steady-state stimulusresponse functions for receptors at the posterior (grey curves; n 20) and
anterior teeth (black curves; n 19), respectively. The solid and
dashed lines represent the mean
1 s.d. of the stimulusresponse
functions. Note the steeper curve at low force levels for the
receptors at anterior teeth indicating a higher sensitivity at low
forces compared with the receptors at posterior teeth [Reprinted
from the Journal of Neurophysiology, Vol. 93, Johnsen SE,
Trulsson M. Encoding of amplitude and rate of tooth loads by
human periodontal afferents from premolar and molar teeth,
pp 18891897, Copyright 2005, with permission from the American Physiological Society].

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M. TRULSSON
steeper average stimulus response curve for receptors at
anterior (grey curve) than at posterior teeth (black
curve). Furthermore, the dynamic sensitivity is both
lower and slower for the posterior receptors. The lower
static and dynamic sensitivity of periodontal receptors
at posterior teeth may reflect a functional adaptation to
the faster and stronger forces that are developed during
motor activities involving the posterior teeth.

Receptor encoding of tooth loads during

biting and chewing
In order to understand how periodontal receptors
contribute to the control of oral motor behaviours such
as biting and chewing, a description of the information
they carry to the brain during these behaviours is
essential. For technical reasons, intraoral recordings are
not easily made from human periodontal afferents
during oral motor functions. Thus, to be able to describe
the responses of human periodontal receptors to arbitrary load profiles, a quantitative model has been
developed that successfully incorporates both the static
and dynamic sensitivity of the receptors (17, 24). In this
model, the discharge rate of a receptor is the weighted
sum of a static response component derived from the
F/(F + c) transform and a dynamic response component
derived from a high-pass filtering of the same trans(a)

(b)

form. The model parameters (weight factors, time

constants of the filter, etc.) are characteristic for each
individual receptor. The results of the modelled discharge rates turned out to be practically identical as
those actually measured for novel force stimulations,
i.e., stimuli that were not included during the development of the model (for further details see Ref. 24).
To predict the receptor discharge evoked by biting
with the anterior teeth, the subject was instructed to
manipulate and bite through a morsel of food resting on
a bar equipped with force transducers (Fig. 7a) (29, 30).
The forces that were developed between the incisors
when cracking a peanut (during a single-phase split
task) and when first holding it briefly between the
teeth (during a two phase hold-and-split task) are
shown in the top panels of Fig. 7b and c, respectively.
Below each force recording is shown the predicted
discharge rates from two periodontal receptors, one
saturating and one non-saturating receptor.
The model predicted that the saturating receptor
would respond distinctly to the small force produced
by the initial contact with the peanut, and would provide
an on-going response when the subject held the peanut
between the teeth (hold phase). Thereby, signalling
information about small changes in force and low levels
of sustained force. However, during rapid exertion of the
higher forces required to split the peanut (split phase),
(c)

Fig. 7. Simulated periodontal receptor responses to empirically recorded force traces (shown at the top) during a split task (b) and a
hold-and-split task (c) with a peanut. The apparatus used to record the force profiles exerted on the food is shown in (a). Subjects were
instructed to position the bar so that the food morsel could be held and split by a pair of opposing central incisors. The food morsel rested
on a horizontal plate of duraluminum equipped with force transducers (FT) for force measurement. Below the force traces in (b) and (c)
are shown simulated receptor responses of a typical saturating and non-saturating anterior periodontal receptor. Note that the two types of
periodontal receptors select out and signal different aspects of the forces generated during the behavioural tasks [Reprinted from Progress
in Neurobiology, Vol. 49, Trulsson M, Johansson RS. Encoding of tooth loads by human periodontal afferents and their role in jaw motor
control, pp 267284, Copyright 1996, with permission from Elsevier].
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SENSORY-MOTOR FUNCTION OF HUMAN PERIODONTAL MECHANORECEPTORS

the receptor exhibited only a moderate and declining
discharge rate. It is noteworthy that the highest discharge rates occurred in response to the initial low force
levels at tooth contact and not to the higher amplitudes
or force rates exerted during the split phase.
In contrast, the non-saturating receptor is predicted
by the model to encode information about force
throughout the split phase. As dynamic sensitivity
parallels static sensitivity, the small forces at the initial
contact with the peanut and during the hold phase
resulted in much lower discharge rates than for the
substantially higher forces generated during the split
phase. However, due to the receptors dynamic sensitivity at high force levels, its discharge rates were
predicted to provide a distorted reflection of the forces
during the split phase. Figure 7c shows that the rate of
force recruitment was momentarily slowed during the
split phase, which resulted in a notch in the discharge
profile rather than two distinct levels of firing.
While the anterior teeth are used during initial food
intake, when morsels are manipulated, split into smaller pieces and transported into the mouth, the posterior
teeth are used during rhythmical chewing when strong
forces are produced by the jaw muscles to grind the
food. During normal chewing of mixed food the forces
exerted on one posterior tooth rarely exceed 5070 N
(31). To predict the discharge from periodontal receptors at posterior teeth evoked during chewing, the

Fig. 8. Simulated periodontal receptor responses to force profiles

recorded during chewing on a force transducer placed between a
pair of molars. Ten different chewing-like force profiles with peak
forces ranging from 3 to 52 N are shown at the top. Below the
force traces, simulated receptor responses are shown for a typical
saturating and a non-saturating posterior periodontal receptor.
Just like the anterior periodontal receptors, the two types of
posterior receptors signal different aspects of the forces acting on
the teeth during normal function [Reprinted from the Journal of
Neurophysiology, Vol. 93, Johnsen SE, Trulsson M. Encoding of
amplitude and rate of tooth loads by human periodontal afferents
from premolar and molar teeth, pp 18891897, Copyright 2005,
with pemission from the American Physiological Society].

subject was instructed to chew on a force transducer

placed between a pair of molars (17). The chewing
forces that were developed between the molars are
shown in the top panel of Fig. 8. Below the force trace
is the predicted discharge rates from two posterior
periodontal receptors, one saturating and one nonsaturating receptor.
The simulations of periodontal receptor responses to
chewing-like force profiles predict that the discharge
rates of the saturating receptors rapidly increase at
initial tooth contact. The receptors continue to discharge as long as the tooth is loaded, but because of the
marked saturation tendencies at higher forces these
receptors poorly encode the magnitude of the strong
chewing forces and the force changes occurring at these
high loads (see the saturating receptor in Fig. 8). The
saturating receptors show their highest discharge rates
shortly after the initial tooth contact and the discharge
rates decrease to a saturated level even if the force
continues to increase. Indeed, response saturation of
the periodontal afferents have been described in
anaesthetized rabbits while chewing on a rubber tube
(32). However, the non-saturating receptors constituted a subpopulation of periodontal receptors that
encoded forces over a wide intensity range (see the
non-saturating receptor in Fig. 8), including forces
developed during the power phase of chewing (31).
Taken together, these observations from the model
simulations demonstrate that periodontal receptors
possess the capacity to signal information about the
mechanical events that occur when humans manipulate and bite food with their anterior teeth and chew
food with their posterior teeth. Receptors with different
force-encoding characteristics capture and emphasize
different aspects of the mechanical events in their
discharge. The simulations suggest that the saturating
and the non-saturating periodontal receptors exhibit
diverse sensory functions during mastication.

Role of periodontal receptors in jaw motor

control
Motor activity for mastication can be generated by
neuronal networks in the brain stem in the absence of
sensory information from the mouth and face (for
review see Ref. 1). However, signals from mechanoreceptors are required for an efficient and adaptive
execution of the masticatory sequence, i.e. from the
acceptance of food to swallowing. As the periodontal

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M. TRULSSON
to biting and provided a means to evaluate this
hypothesis. Three separate observations suggest that
subjects do use periodontal mechanoreceptive information to regulate the level of jaw force during the hold
phase of the hold-and-split task (Fig. 9a). First, the
distribution of hold forces coincide over the range at
which periodontal receptors are most sensitive to
changes in force (Fig. 9c). That is, subjects choose to
use hold forces large enough to achieve a stable clasp
(on average 0.6 N), but automatically avoid higher
forces (>1 N) at which the sensitivity of most receptors
(the saturating receptors) to force changes is compromised. Secondly, during anaesthesia of the periodontal
tissues, the hold forces are considerably increased and
show greater variability, both during the individual
trials and between trials (Fig. 9c; also compare the force
profiles in Fig. 9a and b) (29). Finally, patients lacking
periodontal receptors, such as patients using dental
prostheses supported by the oral mucosa or dental
implants (Fig. 9d) showed similarly high hold force
levels (33). These higher hold forces provided a greater

mechanoreceptors encode information about the temporal, spatial and intensive aspects of tooth loads, it is
likely that their inputs contribute to the regulation of
the muscle activity that generates masticatory forces
and jaw movements. Indeed, there are several investigations, both in animals and man, indicating that
periodontal receptors are involved in the control of
the jaw muscles during biting and chewing (see Refs 1,
2). For a comprehensive account of the inhibitory and
excitatory reflex responses that may be evoked in the
jaw-closing muscles in humans when periodontal
receptors are stimulated, the reader is referred to the
excellent review by Turker (3).
As a large majority of the periodontal receptors
innervating the anterior teeth are highly sensitive to
forces and changes in force levels of <1 N, it was
hypothesized that their input is used for the regulation
of precise manipulative actions involving application of
low forces by the jaw (29). The hold-and-split task
described above simulates the natural situation of
positioning and holding food between the teeth prior
Normal periodontal sensibility

(a)

(b) Anesthesia of the periodontium

Hold-and-split
peanut

Hold-and-split
peanut

10 N

10 N
05 s

30
Normal, hold-and-split trials
Anesthesia, hold-and-split trials

20

20

10

10

0
0

2
Hold force (N)

(d)
4
Hold force (N)

(c)

Static sensitivity imp s1 N1

05 s

Frequency of trials (%)

270

3
2
1
0
Nat Den Imp

Nor Ane

Fig. 9. (a, b) Examples of force profiles (five superimposed trials) obtained during the hold-and-split task with peanuts during normal
periodontal sensibility and during anaesthesia of the periodontium, respectively. Note, the considerably higher and more variable hold
forces produced by the subjects during the periodontal anaesthesia. (c) Frequency distribution of hold forces spontaneously adopted by the
subjects; solid and dashed line histograms refer to trials with normal sensibility and trials with anaesthesia of the periodontium,
respectively. Superimposed curves represent the sensitivity to changes in tooth load of human periodontal afferents. The three dotted
curves refer to the mean
1 s.d. of the first force differential averaged across the 19 periodontal afferents in Fig. 5b. (From Trulsson M,
Johansson RS: Exp Brain Res 107: 486496, 1996.) (d) The left histogram shows the mean hold forces employed by subjects in the natural
(Nat), denture (Den) and implant (Imp) groups. Average values for individual subjects are indicated by filled circles (n 20). Right bar
histograms represent the mean hold force by subjects during normal conditions (Nor) and during periodontal anaesthesia (Ane). Vertical
lines represent the range of average values for individual subjects [Reprinted from the Journal of Dental Research, Trulsson M, Gunne
HSJ. Food-holding and -biting behavior in human subjects lacking periodontal receptors, pp 574-582, Copyright 1998, with permission
from the international and American Associations for Dental Research].
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SENSORY-MOTOR FUNCTION OF HUMAN PERIODONTAL MECHANORECEPTORS

level of security in maintaining the clasp on the morsel
and probably stimulate alternative, less sensitive mechanoreceptors in the tissues that are able to signal its
engagement by the teeth. Furthermore, in anaesthetised subjects, and in patients lacking periodontal
receptors, the morsel frequently escaped from the
incisal edges during the biting task, indicating an
impaired spatial control of the jaw-action vector
(29, 33). Thus, when periodontal afferent information
is lacking, patients show a marked disturbance in the
control of precisely directed, low biting forces, suggesting that periodontal receptors play an important role in
the specification of the level, direction and point of
attack of forces used to hold and manipulate food
between the teeth.
To efficiently handle food during chewing, activation
of the jaw muscles must be coordinated to produce jaw
actions that are spatially adapted to the foods distribution in relation to the teeth. During strong chewing
forces, most of the periodontal receptors are saturated
and poorly encode the magnitude of tooth loads (17).
However, the receptor responses are still dependent on
the direction of the force (10, 24). Thus, during chewing,
the periodontal receptors continuously encode spatial
information about tooth loads and most likely contribute to the spatial control of jaw actions. Indeed, lack of
coordination in chewing has been observed following
denervation of intraoral receptors including periodontal
receptors. The absence of sensory input resulted in
reduced masticatory force and distorted spatial control
of jaw movements during chewing (34, 35).
Given their high sensitivity at low forces, most
periodontal receptors are particularly well suited to
encode in detail the temporal changes in the chewing
force that occur during the early contact phase of each
chewing cycle. From this information, and information
about the movement of the jaw, the mechanical
properties of the food can be derived. Accordingly,
signals from the periodontal receptors may contribute
to the selection of the most appropriate motor signals,
given the existent mechanical food properties. This
would take place in a predictive feed-forward manner
based on learned relationships between patterns of
receptor signals and appropriate efferent signals (cf. 36).
The hypothetical function of early state information
in the control of forthcoming jaw actions brings to mind
that served by tactile receptors in the human fingertips
during grasping and manipulation of small objects.
Immediately following contact with an object, tactile

receptors encode various mechanical contact provisions

including information about the frictional characteristics of the object (36, 37). Such information is used to
trigger the release of motor commands for subsequent
phases of manipulative manoeuvres and to predictively
modify the force output to the physical properties of the
object (38).
These principles are also expressed in the control of
chewing. Ottenhoff et al. (39, 40) have shown that the
additional muscle activity (AMA) required during the
power phase to overcome the impedance of the food is
largely parameterised in advance on the basis of sensory
experiences during the preceding chewing cycle. Memory of the foods resistance in the previous cycle is used
to predictively scale the muscle commands that initiate
the power phase. It is also used to regulate the strength
of a sensory mediated reflex increase in muscle
activity elicited by contact with the food resistance; a
maximum output strength of the reflex contribution
to the AMA is set (40). Signals from periodontal
receptors may contribute both to information about
the food resistance gained during the previous cycle,
and to the reflex mediated component of the AMA
(40).
Studies on anaesthetized rabbits performing rhythmical jaw movements confirm that the AMA often
precedes the tooth contact, supporting the operation of
a feed-forward control mechanism (41). Blocking of the
periodontal receptor input did not affect the AMA,
whereas blocking of information from muscle spindle
receptors innervating the jaw-closing muscles abolished
the AMA preceding the tooth contact. Earlier animal
studies have shown that combined destruction of
periodontal receptors and muscle-spindle receptors
diminish the AMA almost completely, while destruction of either one of both receptors do not completely
diminish the AMA (34, 35, 42). These results suggest
that the timing of the predictive component of the
AMA may be controlled mainly by information from
muscle spindles, while the magnitude of the AMA is
under the control of both muscle spindles and periodontal receptors (41). Furthermore, blocking of information from periodontal receptors in anaesthetized
rabbits significantly reduces the build-up speed of the
masticatory force during chewing (43). Thus, information acquired by periodontal receptors early after
contact during each chewing cycle may be used to
scale the muscle commands of the ensuing power phase
to the current food impedance. Indeed, jaw actions

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M. TRULSSON
observably adapt to the gradually changing mechanical
properties of a bolus also during natural chewing (34,
4446). Interestingly, in contrast to subjects with their
natural teeth, patients with prostheses supported by
dental implants seem to chew with about the same
pattern of muscle activity throughout the entire masticatory sequence (47). The lack of adaptation in these
patients can be reasonably attributed to the absence of
periodontal receptors.
In summary, signals from periodontal receptors are
used in the fine motor control of jaw actions associated
with biting, intraoral manipulation and the chewing of
food. It is clear from studies of various patient groups
(e.g. patients with dental implants) that important
sensory-motor functions are lost or impaired when
these receptors are removed during the extraction of
teeth. This knowledge emphasizes the importance of
maintaining natural teeth with healthy periodontal
function whenever possible.

10.

11.

12.

13.
14.

15.

16.

17.

Acknowledgments
This work was supported by the Karolinska Institute
and the Swedish Medical Research Council.

18.

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Correspondence: Dr M. Trulsson, Institute of Odontology, Karolinska

Institutet, Box 4064, S-141 04 Huddinge, Sweden.
E-mail: mats.trulsson@ki.se

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